Localised Sensitiveness to Light, and Its Transmitted Effects

Phalaris Canariensis.—Whilst observing the accuracy with which the cotyledons of this plant became bent towards the light of a small lamp, we were impressed with the idea that the uppermost part determined the direction of the curvature of the lower part. When the cotyledons are exposed to a lateral light, the upper part bends first, and afterwards the bending gradually extends down to the base, and, as we shall presently see, even a little beneath the ground. This holds good with cotyledons from less than .1 inch (one was observed to act in this manner which was only .03 in height) to about .5 of an inch in height; but when they have grown to nearly an inch in height, the basal part, for a length of .15 to .2 of an inch above the ground, ceases to bend. As with young cotyledons the lower part goes on bending, after the upper part has become well arched towards a lateral light, the apex would ultimately point to the ground instead of to the light, did not the upper part reverse its curvature and straighten itself, as soon as the upper convex surface of the bowed-down portion received more light than the lower concave surface. The position ultimately assumed by young and upright cotyledons, exposed to light entering obliquely from above through a window, is shown in the accompanying figure (Fig. 181); and here it may be seen that the whole upper part has become very nearly straight. When the cotyledons were exposed before a bright lamp, standing on the same level with them, the upper part, which was at first greatly arched towards the light, became straight and strictly parallel with the surface of the soil in the pots; the basal part being now rectangularly bent. All this great amount of curvature, together with the subsequent straightening of the upper part, was often effected in a few hours.

Fig. 181. Phalaris Canariensis: cotyledons after exposure in a box open on one side in front of a south-west window during 8 h. Curvature towards the light accurately traced. The short horizontal lines show the level of the ground.

After the uppermost part has become bowed a little to the light, its overhanging weight must tend to increase the curvature of the lower part; but any such effect was shown in several ways to be quite insignificant. When little caps of tin-foil (hereafter to be described) were placed on the summits of the cotyledons, though this must have added considerably to their weight, the rate or amount of bending was not thus increased. But the best evidence was afforded by placing pots with seedlings of Phalaris before a lamp in such a position, that the cotyledons were horizontally extended and projected at right angles to the line of light. In the course of 5½ h. they were directed towards the light with their bases bent at right angles; and this abrupt curvature could not have been aided in the least by the weight of the upper part, which acted at right angles to the plane of curvature.

It will be shown that when the upper halves of the cotyledons of Phalaris and Avena were enclosed in little pipes of tin-foil or of blackened glass, in which case the upper part was mechanically prevented from bending, the lower and unenclosed part did not bend when exposed to a lateral light; and it occurred to us that this fact might be due, not to the exclusion of the light from the upper part, but to some necessity of the bending gradually travelling down the cotyledons, so that unless the upper part first became bent, the lower could not bend, however much it might be stimulated. It was necessary for our purpose to ascertain whether this notion was true, and it was proved false; for the lower halves of several cotyledons became bowed to the light, although their upper halves were enclosed in little glass tubes (not blackened), which prevented, as far as we could judge, their bending. Nevertheless, as the part within the tube might possibly bend a very little, fine rigid rods or flat splinters of thin glass were cemented with shellac to one side of the upper part of 15 cotyledons; and in six cases they were in addition tied on with threads. They were thus forced to remain quite straight. The result was that the lower halves of all became bowed to the light, but generally not in so great a degree as the corresponding part of the free seedlings in the same pots; and this may perhaps be accounted for by some slight degree of injury having been caused by a considerable surface having been smeared with shellac. It may be added, that when the cotyledons of Phalaris and Avena are acted on by apogeotropism, it is the upper part which begins first to bend; and when this part was rendered rigid in the manner just described, the upward curvature of the basal part was not thus prevented.

To test our belief that the upper part of the cotyledons of Phalaris, when exposed to a lateral light, regulates the bending of the lower part, many experiments were tried; but most of our first attempts proved useless from various causes not worth specifying. Seven cotyledons had their tips cut off for lengths varying between .1 and .16 of an inch, and these, when left exposed all day to a lateral light, remained upright. In another set of 7 cotyledons, the tips were cut off for a length of only about .05 of an inch (1.27 mm.) and these became bowed towards a lateral light, but not nearly so much as the many other seedlings in the same pots. This latter case shows that cutting off the tips does not by itself injure the plants so seriously as to prevent heliotropism; but we thought at the time, that such injury might follow when a greater length was cut off, as in the first set of experiments. Therefore, no more trials of this kind were made, which we now regret; as we afterwards found that when the tips of three cotyledons were cut off for a length of .2 inch, and of four others for lengths of .14, .12, .1, and .07 inch, and they were extended horizontally, the amputation did not interfere in the least with their bending vertically upwards, through the action of apogeotropism, like unmutilated specimens. It is therefore extremely improbable that the amputation of the tips for lengths of from .1 to .14 inch, could from the injury thus caused have prevented the lower part from bending towards the light.

We next tried the effects of covering the upper part of the cotyledons of Phalaris with little caps which were impermeable to light; the whole lower part being left fully exposed before a south-west window or a bright paraffin lamp. Some of the caps were made of extremely thin tin-foil blackened within; these had the disadvantage of occasionally, though rarely, being too heavy, especially when twice folded. The basal edges could be pressed into close contact with the cotyledons; though this again required care to prevent injuring them. Nevertheless, any injury thus caused could be detected by removing the caps, and trying whether the cotyledons were then sensitive to light. Other caps were made of tubes of the thinnest glass, which when painted black served well, with the one great disadvantage that the lower ends could not be closed. But tubes were used which fitted the cotyledons almost closely, and black paper was placed on the soil round each, to check the upward reflection of light from the soil. Such tubes were in one respect far better than caps of tin-foil, as it was possible to cover at the same time some cotyledons with transparent and others with opaque tubes; and thus our experiments could be controlled. It should be kept in mind that young cotyledons were selected for trial, and that these when not interfered with become bowed down to the ground towards the light.

We will begin with the glass-tubes. The summits of nine cotyledons, differing somewhat in height, were enclosed for rather less than half their lengths in uncoloured or transparent tubes; and these were then exposed before a south-west window on a bright day for 8 h. All of them became strongly curved towards the light, in the same degree as the many other free seedlings in the same pots; so that the glass-tubes certainly did not prevent the cotyledons from bending towards the light. Nineteen other cotyledons were, at the same time, similarly enclosed in tubes thickly painted with Indian ink. On five of them, the paint, to our surprise, contracted after exposure to the sunlight, and very narrow cracks were formed, through which a little light entered; and these five cases were rejected. Of the remaining 14 cotyledons, the lower halves of which had been fully exposed to the light for the whole time, 7 continued quite straight and upright; 1 was considerably bowed to the light, and 6 were slightly bowed, but with the exposed bases of most of them almost or quite straight. It is possible that some light may have been reflected upwards from the soil and entered the bases of these 7 tubes, as the sun shone brightly, though bits of blackened paper had been placed on the soil round them. Nevertheless, the 7 cotyledons which were slightly bowed, together with the 7 upright ones, presented a most remarkable contrast in appearance with the many other seedlings in the same pots to which nothing had been done. The blackened tubes were then removed from 10 of these seedlings, and they were now exposed before a lamp for 8 h.; 9 of them became greatly, and 1 moderately, curved towards the light, proving that the previous absence of any curvature in the basal part, or the presence of only a slight degree of curvature there, was due to the exclusion of light from the upper part.

Similar observations were made on 12 younger cotyledons with their upper halves enclosed within glass-tubes coated with black varnish, and with their lower halves fully exposed to bright sunshine. In these younger seedlings the sensitive zone seems to extend rather lower down, as was observed on some other occasions, for two became almost as much curved towards the light as the free seedlings; and the remaining ten were slightly curved, although the basal part of several of them, which normally becomes more curved than any other part, exhibited hardly a trace of curvature. These 12 seedlings taken together differed greatly in their degree of curvature from all the many other seedlings in the same pots.

Better evidence of the efficiency of the blackened tubes was incidentally afforded by some experiments hereafter to be given, in which the upper halves of 14 cotyledons were enclosed in tubes from which an extremely narrow stripe of the black varnish had been scraped off. These cleared stripes were not directed towards the window, but obliquely to one side of the room, so that only a very little light could act on the upper halves of the cotyledons. These 14 seedlings remained during eight hours of exposure before a south-west window on a hazy day quite upright; whereas all the other many free seedlings in the same pots became greatly bowed towards the light.

We will now turn to the trials with caps made of very thin tin-foil. These were placed at different times on the summits of 24 cotyledons, and they extended down for a length of between .15 and .2 of an inch. The seedlings were exposed to a lateral light for periods varying between 6 h. 30 m. and 7 h. 45 m., which sufficed to cause all the other seedlings in the same pots to become almost rectangularly bent towards the light. They varied in height from only .04 to 1.15 inch, but the greater number were about .75 inch. Of the 24 cotyledons with their summits thus protected, 3 became much bent, but not in the direction of the light, and as they did not straighten themselves through apogeotropism during the following night, either the caps were too heavy or the plants themselves were in a weak condition; and these three cases may be excluded. There are left for consideration 21 cotyledons; of these 17 remained all the time quite upright; the other 4 became slightly inclined to the light, but not in a degree comparable with that of the many free seedlings in the same pots. As the glass-tubes, when unpainted, did not prevent the cotyledons from becoming greatly bowed, it cannot be supposed that the caps of very thin tin-foil did so, except through the exclusion of the light. To prove that the plants had not been injured, the caps were removed from 6 of the upright seedlings, and these were exposed before a paraffin lamp for the same length of time as before, and they now all became greatly curved towards the light.

As caps between .15 and .2 of an inch in depth were thus proved to be highly efficient in preventing the cotyledons from bending towards the light, 8 other cotyledons were protected with caps between only .06 and .12 in depth. Of these, two remained vertical, one was considerably and five slightly curved towards the light, but far less so than the free seedlings in the same pots.

Another trial was made in a different manner, namely, by bandaging with strips of tin-foil, about .2 in breadth, the upper part, but not the actual summit, of eight moderately young seedlings a little over half an inch in height. The summits and the basal parts were thus left fully exposed to a lateral light during 8 h.; an upper intermediate zone being protected. With four of these seedlings the summits were exposed for a length of .05 inch, and in two of them this part became curved towards the light, but the whole lower part remained quite upright; whereas the entire length of the other two seedlings became slightly curved towards the light. The summits of the four other seedlings were exposed for a length of .04 inch, and of these one remained almost upright, whilst the other three became considerably curved towards the light. The many free seedlings in the same pots were all greatly curved towards the light.

From these several sets of experiments, including those with the glass-tubes, and those when the tips were cut off, we may infer that the exclusion of light from the upper part of the cotyledons of Phalaris prevents the lower part, though fully exposed to a lateral light, from becoming curved. The summit for a length of .04 or .05 of an inch, though it is itself sensitive and curves towards the light, has only a slight power of causing the lower part to bend. Nor has the exclusion of light from the summit for a length of .1 of an inch a strong influence on the curvature of the lower part. On the other hand, an exclusion for a length of between .15 and .2 of an inch, or of the whole upper half, plainly prevents the lower and fully illuminated part from becoming curved in the manner (see Fig. 181) which invariably occurs when a free cotyledon is exposed to a lateral light. With very young seedlings the sensitive zone seems to extend rather lower down relatively to their height than in older seedlings. We must therefore conclude that when seedlings are freely exposed to a lateral light some influence is transmitted from the upper to the lower part, causing the latter to bend.

This conclusion is supported by what may be seen to occur on a small scale, especially with young cotyledons, without any artificial exclusion of the light; for they bend beneath the earth where no light can enter. Seeds of Phalaris were covered with a layer one-fourth of an inch in thickness of very fine sand, consisting of extremely minute grains of silex coated with oxide of iron. A layer of this sand, moistened to the same degree as that over the seeds, was spread over a glass-plate; and when the layer was .05 of an inch in thickness (carefully measured) no light from a bright sky could be seen to pass through it, unless it was viewed through a long blackened tube, and then a trace of light could be detected, but probably much too little to affect any plant. A layer .1 of an inch in thickness was quite impermeable to light, as judged by the eye aided by the tube. It may be worth adding that the layer, when dried, remained equally impermeable to light. This sand yielded to very slight pressure whilst kept moist, and in this state did not contract or crack in the least. In a first trial, cotyledons which had grown to a moderate height were exposed for 8 h. before a paraffin lamp, and they became greatly bowed. At their bases on the shaded side opposite to the light, well-defined, crescentic, open furrows were formed, which (measured under a microscope with a micrometer) were from .02 to .03 of an inch in breadth, and these had evidently been left by the bending of the buried bases of the cotyledons towards the light. On the side of the light the cotyledons were in close contact with the sand, which was a very little heaped up. By removing with a sharp knife the sand on one side of the cotyledons in the line of the light, the bent portion and the open furrows were found to extend down to a depth of about .1 of an inch, where no light could enter. The chords of the short buried arcs formed in four cases angles of 11°, 13°, 15°, and 18°, with the perpendicular. By the following morning these short bowed portions had straightened themselves through apogeotropism.

In the next trial much younger cotyledons were similarly treated, but were exposed to a rather obscure lateral light. After some hours, a bowed cotyledon, .3 inch in height, had an open furrow on the shaded side .04 inch in breadth; another cotyledon, only .13 inch in height, had left a furrow .02 inch in breadth. But the most curious case was that of a cotyledon which had just protruded above the ground and was only .03 inch in height, and this was found to be bowed in the direction of the light to a depth of .2 of an inch beneath the surface. From what we know of the impermeability of this sand to light, the upper illuminated part in these several cases must have determined the curvature of the lower buried portions. But an apparent cause of doubt may be suggested: as the cotyledons are continually circumnutating, they tend to form a minute crack or furrow all round their bases, which would admit a little light on all sides; but this would not happen when they were illuminated laterally, for we know that they quickly bend towards a lateral light, and they then press so firmly against the sand on the illuminated side as to furrow it, and this would effectually exclude light on this side. Any light admitted on the opposite and shaded side, where an open furrow is formed, would tend to counteract the curvature towards the lamp or other source of the light. It may be added, that the use of fine moist sand, which yields easily to pressure, was indispensable in the above experiments; for seedlings raised in common soil, not kept especially damp, and exposed for 9 h. 30 m. to a strong lateral light, did not form an open furrow at their bases on the shaded side, and were not bowed beneath the surface.

Perhaps the most striking proof of the action of the upper on the lower part of the cotyledons of Phalaris, when laterally illuminated, was afforded by the blackened glass-tubes (before alluded to) with very narrow stripes of the varnish scraped off on one side, through which a little light was admitted. The breadth of these stripes or slits varied between .01 and .02 inch (.25 and .51 mm.). Cotyledons with their upper halves enclosed in such tubes were placed before a south-west window, in such a position, that the scraped stripes did not directly face the window, but obliquely to one side. The seedlings were left exposed for 8 h., before the close of which time the many free seedlings in the same pots had become greatly bowed towards the window. Under these circumstances, the whole lower halves of the cotyledons, which had their summits enclosed in the tubes, were fully exposed to the light of the sky, whilst their upper halves received exclusively or chiefly diffused light from the room, and this only through a very narrow slit on one side. Now, if the curvature of the lower part had been determined by the illumination of this part, all the cotyledons assuredly would have become curved towards the window; but this was far from being the case. Tubes of the kind just described were placed on several occasions over the upper halves of 27 cotyledons; 14 of them remained all the time quite vertical; so that sufficient diffused light did not enter through the narrow slits to produce any effect whatever; and they behaved in the same manner as if their upper halves had been enclosed in completely blackened tubes. The lower halves of the 13 other cotyledons became bowed not directly in the line of the window, but obliquely towards it; one pointed at an angle of only 18°, but the remaining 12 at angles varying between 45° and 62° from the line of the window. At the commencement of the experiment, pins had been laid on the earth in the direction towards which the slits in the varnish faced; and in this direction alone a small amount of diffused light entered. At the close of the experiment, 7 of the bowed cotyledons pointed exactly in the line of the pins, and 6 of them in a line between that of the pins and that of the window. This intermediate position is intelligible, for any light from the sky which entered obliquely through the slits would be much more efficient than the diffused light which entered directly through them. After the 8 h. exposure, the contrast in appearance between these 13 cotyledons and the many other seedlings in the same pots, which were all (excepting the above 14 vertical ones) greatly bowed in straight and parallel lines towards the window, was extremely remarkable. It is therefore certain that a little weak light striking the upper halves of the cotyledons of Phalaris, is far more potent in determining the direction of the curvature of the lower halves, than the full illumination of the latter during the whole time of exposure.

In confirmation of the above results, the effect of thickly painting with Indian ink one side of the upper part of three cotyledons of Phalaris, for a length of .2 inch from their tips, may be worth giving. These were placed so that the unpainted surface was directed not towards the window, but a little to one side; and they all became bent towards the unpainted side, and from the line of the window by angles amounting to 31°, 35°, and 83°. The curvature in this direction extended down to their bases, although the whole lower part was fully exposed to the light from the window.

Finally, although there can be no doubt that the illumination of the upper part of the cotyledons of Phalaris greatly affects the power and manner of bending of the lower part, yet some observations seemed to render it probable that the simultaneous stimulation of the lower part by light greatly favours, or is almost necessary, for its well-marked curvature; but our experiments were not conclusive, owing to the difficulty of excluding light from the lower halves without mechanically preventing their curvature.

Avena sativa.—The cotyledons of this plant become quickly bowed towards a lateral light, exactly like those of Phalaris. Experiments similar to the foregoing ones were tried, and we will give the results as briefly as possible. They are somewhat less conclusive than in the case of Phalaris, and this may possibly be accounted for by the sensitive zone varying in extension, in a species so long cultivated and variable as the common Oat. Cotyledons a little under three-quarters of an inch in height were selected for trial: six had their summits protected from light by tin-foil caps, .25 inch in depth, and two others by caps .3 inch in depth. Of these 8 cotyledons, five remained upright during 8 hours of exposure, although their lower parts were fully exposed to the light all the time; two were very slightly, and one considerably, bowed towards it. Caps only .2 or .22 inch in depth were placed over 4 other cotyledons, and now only one remained upright, one was slightly, and two considerably bowed to the light. In this and the following cases all the free seedlings in the same pots became greatly bowed to the light.

Our next trial was made with short lengths of thin and fairly transparent quills; for glass-tubes of sufficient diameter to go over the cotyledons would have been too heavy. Firstly, the summits of 13 cotyledons were enclosed in unpainted quills, and of these 11 became greatly and 2 slightly bowed to the light; so that the mere act of enclosure did not prevent the lower part from becoming bowed. Secondly, the summits of 11 cotyledons were enclosed in quills .3 inch in length, painted so as to be impermeable to light; of these, 7 did not become at all inclined towards the light, but 3 of them were slightly bent more or less transversely with respect to the line of light, and these might perhaps have been altogether excluded; one alone was slightly bowed towards the light. Painted quills, .25 inch in length, were placed over the summits of 4 other cotyledons; of these, one alone remained upright, a second was slightly bowed, and the two others as much bowed to the light as the free seedlings in the same pots. These two latter cases, considering that the caps were .25 in length, are inexplicable.

Lastly, the summits of 8 cotyledons were coated with flexible and highly transparent gold-beaters’ skin, and all became as much bowed to the light as the free seedlings. The summits of 9 other cotyledons were similarly coated with gold-beaters’ skin, which was then painted to a depth of between .25 and .3 inch, so as to be impermeable to light; of these 5 remained upright, and 4 were well bowed to the light, almost or quite as well as the free seedlings. These latter four cases, as well as the two in the last paragraph, offer a strong exception to the rule that the illumination of the upper part determines the curvature of the lower part. Nevertheless, 5 of these 8 cotyledons remained quite upright, although their lower halves were fully illuminated all the time; and it would almost be a prodigy to find five free seedlings standing vertically after an exposure for several hours to a lateral light.

The cotyledons of Avena, like those of Phalaris, when growing in soft, damp, fine sand, leave an open crescentric furrow on the shaded side, after bending to a lateral light; and they become bowed beneath the surface at a depth to which, as we know, light cannot penetrate. The arcs of the chords of the buried bowed portions formed in two cases angles of 20° and 21° with the perpendicular. The open furrows on the shaded side were, in four cases, .008, .016, .024, and .024 of an inch in breadth. Brassica oleracea (Common Red).—It will here be shown that the upper half of the hypocotyl of the cabbage, when illuminated by a lateral light, determines the curvature of the lower half. It is necessary to experimentise on young seedlings about half an inch or rather less in height, for when grown to an inch and upwards the basal part ceases to bend. We first tried painting the hypocotyls with Indian ink, or cutting off their summits for various lengths; but these experiments are not worth giving, though they confirm, as far as they can be trusted, the results of the following ones. These were made by folding gold-beaters’ skin once round the upper halves of young hypocotyls, and painting it thickly with Indian ink or with black grease. As a control experiment, the same transparent skin, left unpainted, was folded round the upper halves of 12 hypocotyls; and these all became greatly curved to the light, excepting one, which was only moderately curved. Twenty other young hypocotyls had the skin round their upper halves painted, whilst their lower halves were left quite uncovered. These seedlings were then exposed, generally for between 7 and 8 h., in a box blackened within and open in front, either before a south-west window or a paraffin lamp. This exposure was amply sufficient, as was shown by the strongly-marked heliotropism of all the free seedlings in the same pots; nevertheless, some were left exposed to the light for a much longer time. Of the 20 hypocotyls thus treated, 14 remained quite upright, and 6 became slightly bowed to the light; but 2 of these latter cases were not really exceptions, for on removing the skin the paint was found imperfect and was penetrated by many small transparent spaces on the side which faced the light. Moreover, in two other cases the painted skin did not extend quite halfway down the hypocotyl. Although there was a wonderful contrast in the several pots between these 20 hypocotyls and the other many free seedlings, which were all greatly bowed down to their bases in the direction of the light, some being almost prostrate on the ground.

The most successful trial on any one day (included in the above results) is worth describing in detail. Six young seedlings were selected, the hypocotyls of which were nearly .45 inch, excepting one, which was .6 inch in height, measured from the bases of their petioles to the ground. Their upper halves, judged as accurately as could be done by the eye, were folded once round with gold-beaters’ skin, and this was painted thickly with Indian ink. They were exposed in an otherwise darkened room before a bright paraffin lamp, which stood on a level with the two pots containing the seedlings. They were first looked at after an interval of 5 h. 10 m., and five of the protected hypocotyls were found quite erect, the sixth being very slightly inclined to the light; whereas all the many free seedlings in the same two pots were greatly bowed to the light. They were again examined after a continuous exposure to the light of 20 h. 35m.; and now the contrast between the two sets was wonderfully great; for the free seedlings had their hypocotyls extended almost horizontally in the direction of the light, and were curved down to the ground; whilst those with the upper halves protected by the painted skin, but with their lower halves fully exposed to the light, still remained quite upright, with the exception of the one which retained the same slight inclination to the light which it had before. This latter seedling was found to have been rather badly painted, for on the side facing the light the red colour of the hypocotyl could be distinguished through the paint.

We next tried nine older seedlings, the hypocotyls of which varied between 1 and 1.6 inch in height. the gold-beaters’ skin round their upper parts was painted with black grease to a depth of only .3 inch, that is, from less than a third to a fourth or fifth of their total heights. They were exposed to the light for 7 h. 15 m.; and the result showed that the whole of the sensitive zone, which determines the curvature of the lower part, was not protected from the action of the light; for all 9 became curved towards it, 4 of them very slightly, 3 moderately, and 2 almost as much as the unprotected seedlings. Nevertheless, the whole 9 taken together differed plainly in their degree of curvature from the many free seedlings, and from some which were wrapped in unpainted skin, growing in the same two pots.

Seeds were covered with about a quarter of an inch of the fine sand described under Phalaris; and when the hypocotyls had grown to a height of between .4 and .55 inch, they were exposed during 9 h. before a paraffin lamp, their bases being at first closely surrounded by the damp sand. They all became bowed down to the ground, so that their upper parts lay near to and almost parallel to the surface of the soil. On the side of the light their bases were in close contact with the sand, which was here a very little heaped up; on the opposite or shaded side there were open, crescentic cracks or furrows, rather above .01 of an inch in width; but they were not so sharp and regular as those made by Phalaris and Avena, and therefore could not be so easily measured under the microscope. The hypocotyls were found, when the sand was removed on one side, to be curved to a depth beneath the surface in three cases of at least .1 inch, in a fourth case of .11, and in a fifth of .15 inch. The chords of the arcs of the short, buried, bowed portions formed angles of between 11° and 15° with the perpendicular. From what we have seen of the impermeability of this sand to light, the curvature of the hypocotyls certainly extended down to a depth where no light could enter; and the curvature must have been caused by an influence transmitted from the upper illuminated part.

The lower halves of five young hypocotyls were surrounded by unpainted gold-beaters’ skin, and these, after an exposure of 8 h. before a paraffin lamp, all became as much bowed to the light as the free seedlings. The lower halves of 10 other young hypocotyls, similarly surrounded with the skin, were thickly painted with Indian ink; their upper and unprotected halves became well curved to the light, but their lower and protected halves remained vertical in all the cases excepting one, and on this the layer of paint was imperfect. This result seems to prove that the influence transmitted from the upper part is not sufficient to cause the lower part to bend, unless it be at the same time illuminated; but there remains the doubt, as in the case of Phalaris, whether the skin covered with a rather thick crust of dry Indian ink did not mechanically prevent their curvature.

Beta vulgaris.—A few analogous experiments were tried on this plant, which is not very well adapted for the purpose, as the basal part of the hypocotyl, after it has grown to above half an inch in height, does not bend much on exposure to a lateral light. Four hypocotyls were surrounded close beneath their petioles with strips of thin tin-foil, .2 inch in breadth, and they remained upright all day before a paraffin lamp; two others were surrounded with strips .15 inch in breadth, and one of these remained upright, the other becoming bowed; the bandages in two other cases were only .1 inch in breadth, and both of these hypocotyls became bowed, though one only slightly, towards the light. The free seedlings in the same pots were all fairly well curved towards the light; and during the following night became nearly upright. The pots were now turned round and placed before a window, so that the opposite sides of the seedlings were exposed to the light, towards which all the unprotected hypocotyls became bent in the course of 7 h. Seven out of the 8 seedlings with bandages of tin-foil remained upright, but one which had a bandage only .1 inch in breadth, became curved to the light. On another occasion, the upper halves of 7 hypocotyls were surrounded with painted gold-beaters’ skin; of these 4 remained upright, and 3 became a little curved to the light: at the same time 4 other seedlings surrounded with unpainted skin, as well as the free ones in the same pots, all became bowed towards the lamp, before which they had been exposed during 22 hours.

Radicles of Sinapis alba.—The radicles of some plants are indifferent, as far as curvature is concerned, to the action of light; whilst others bend towards and others from it. Whether these movements are of any service to the plant is very doubtful, at least in the case of subterranean roots; they probably result from the radicles being sensitive to contact, moisture, and gravitation, and as a consequence to other irritants which are never naturally encountered. The radicles of Sinapis alba, when immersed in water and exposed to a lateral light, bend from it, or are apheliotropic. They become bent for a length of about 4 mm. from their tips. To ascertain whether this movement generally occurred, 41 radicles, which had germinated in damp sawdust, were immersed in water and exposed to a lateral light; and they all, with two doubtful exceptions, became curved from the light. At the same time the tips of 54 other radicles, similarly exposed, were just touched with nitrate of silver. They were blackened for a length of from .05 to .07 mm., and probably killed; but it should be observed that this did not check materially, if at all, the growth of the upper part; for several, which were measured, increased in the course of only 8–9 h. by 5 to 7 mm. in length. Of the 54 cauterised radicles one case was doubtful, 25 curved themselves from the light in the normal manner, and 28, or more than half, were not in the least apheliotropic. There was a considerable difference, which we cannot account for, in the results of the experiments tried towards the end of April and in the middle of September. Fifteen radicles (part of the above 54) were cauterised at the former period and were exposed to sunshine, of which 12 failed to be apheliotropic, 2 were still apheliotropic, and 1 was doubtful. In September, 39 cauterised radicles were exposed to a northern light, being kept at a proper temperature; and now 23 continued to be apheliotropic in the normal manner, and only 16 failed to bend from the light. Looking at the aggregate results at both periods, there can be no doubt that the destruction of the tip for less than a millimeter in length destroyed in more than half the cases their power of moving from the light. It is probable that if the tips had been cauterised for the length of a whole millimeter, all signs of apheliotropism would have disappeared. It may be suggested that although the application of caustic does not stop growth, yet enough may be absorbed to destroy the power of movement in the upper part; but this suggestion must be rejected, for we have seen and shall again see, that cauterising one side of the tip of various kinds of radicles actually excites movement. The conclusion seems inevitable that sensitiveness to light resides in the tip of the radicle of Sinapis alba; and that the tip when thus stimulated transmits some influence to the upper part, causing it to bend. The case in this respect is parallel with that of the radicles of several plants, the tips of which are sensitive to contact and to other irritants, and, as will be shown in the eleventh chapter, to gravitation.